sign in sign up
<<
Home , Atheris, Atheris chlorechis

Volume 12, Number 2 April2002 ISSN 0268-0130 THE HERPETOLOGICAL JOURNAL

Published by the Indexed in BRITISH HERPETOLOGICAL SOCIETY Current Contents HERPETOLOGJCAL JOURNAL, Vol. 12, pp. 55-6 1 (2002)

A NEW (SERPENTES: ), FROM TAI NATIONAL PARK,

R . ERNST AND M.-0. RODEL

Th eodor Boveri Institute Biocenter of the University, Department of Ecology and Trop ical Biology (Zoology /II) . Am Hub/and, D-97074 Wiirzburg, Germany

We describe a new species of the Atheris fromTa'i National Park, a large area in south-western Ivory Coast. Atheris sp. nov. shows close affinities to A. squamigera, but is distinguished from it by a combination of scale characteristics, as well as morphometric diffe rences in head proportions.

Key words: Serpentes, Yiperidae, Atlzeris sp. nov., Ta'i National Park, Ivory Coast;

INTRODUCTION The geographical co-ordinates were obtained from The genus Atheris has undergone several taxonomic Geographic Names Processing System - Phase IV, and a revisions (Broadley, 1996, 1998). However, due to the portable GPS (Garmin 12XL). The map was drawn with huge variability within the genus (Jacobi, 2001 ), the the mapping program Versamap version 2.07. taxonomic status of several taxa still remains to be set­ In order to ensure comparability, we investigated the tled. Very recently two new species of A theris have been same characters used in recent treatments of the genus described: A. acuminata from (Broadley, 1998) (Broadley, 1998; Lawson & Ustach, 2000): and A. broadleyi from (Lawson, 1999). The suprarostrals (SRO), internasals (INS), interorbitals resurrection of A. subocularis from the synonymy of A. (IOS), maximum transverse head scales (MTHS), squamigera has currently been published (Lawson, circumorbital scales (COS), interoculabials (IOL), Noonan & Ustach, 2001). The latter species is espe­ interocunasals (ION), supralabials (SL), infralabials cially renowned forits variability (Spawls & Branch, (IL), pairs of sublinguals (PSL), dorsal scale rows at 1995). However, previous papers may have inadvert­ mid-body (MSR), ventrals and subcaudals. For discus­ ently increased its known variation in many characters sion of the diagnostic significance of these characters, by combining distinct species (Lawson, 1999). see Broadley (1998). In addition we investigated According to Hughes (pers. comm.), without a thor­ morphometric proportions of the head, as well as struc­ ough re-examination of existing museum specimens, it ture and shape of scales. Scale terminology is according cannot be decided whether A. squamigera is a geo­ to Broadley ( 1998). graphically variable species or a species complex. Head width (HW) was always taken as the maximum However, there is good evidence of more than one spe­ head width possible. Head length (HL) was measured cies being involved. Atheris hirsuta sp. nov. from Ta'i from snout to the quadratum. Interorbital distance (IOD) National Park (TNP), Ivory Coast, is one of these spe­ was measured between the circumorbital scale rows. cies that shows close affinities to A. squamigera, but Snout-eye distance (SED) was measured from the ante­ differs sufficiently to be recognized at the species level. rior comer of the eye to the snout tip. Scale and head measures were taken to the nearest 0.1 mm using a dial MATERIAL AND METHODS calliper. When scale counts differed between the sides of the head, we gave both values. Snout-vent length For comparison, we investigated museum specimens (SVL) was taken to the nearest mm using a metre rule. of A. squamigera fromthe entire Central and West Afri­ Results are summarized in Tables 1, 2 and 3. Scale can range of the species (Fig. 1). Museum specimens microstructure was examined by means of a scanning under investigation originated from, and are deposited electron microscope (SEM). in, the following collections: Staatliches Museum fur Naturkunde Stuttgart (SMNS), Zoologisches SPECIES ACCOUNT Forschungsinstitut und Museum Alexander Koenig ATHERIS HJRSUTA SP. NOV. Bonn (ZFMK), Zoologisches Museum der Humboldt Universitiitzu Berlin (ZMB), Zoologisches Institut und Holotype. SMNS 11333, male, about 6 km West of Zoologisches Museum der Universitiit Hamburg the "Station de Recherche en Ecologie Tropicale" (ZMH), Zoological Museum, University of Copenhagen (SRET 5°50'N 7°19'W), Ta'iNational Park, Ivory Coast, (ZMUC), and Zoologische Staatssammlung Mtinchen 20 September 2000, 07.00 hrs, R. Ernst leg. (ZSM). Diagnosis. Slender tree viper with rather short head, IODISED 2.3; heavily carinated scales, especially on head and neck, giving the a bristly appearance; Correspondence: R. Ernst, Theodor Boveri Institute Biocenter of the University, Department of Animal Ecology keels run in long curves towards a sharp tip; six and Tropical Biology (Zoology III), Am Hubland, D-97074 suprarostrals; eight to nine infralabials; three pairs of Wi.irzburg, Germany. sublinguals; elongate dorsal scales; 16 scale rows E-mail: [email protected] around mid-body. 56 R. ERNST ET AL.

w I0° E 30° interorbitals; 14 scales between posterior supralabials;

eye large, 3.1 mm in diameter; eye diameter 2.5. times the eye-border to lip distance; no interoculabial; 14 and 15 circumorbitals, fo ur in contact with supralabials; nine and 10 supralabials; mental twice as wide as long; eight and nine infralabials,first pair in contact posterior to mental; three pairs of sublinguals; three preventrals; two to three rows of keeled gular scales; dorsal scales heavily keeled, especially on head and anterior third of body; keels run curved to a sharp tip; lateral scales with­ out serration; no reticulate microdermatoglyphicpattern FIG. I. West and Central African distribution of A. of cell boundary lines; size of the scale keels decreases squamigera (circles) and type-locality ofA. hirsuta sp. nov. from head to tail; keels at mid-body shorter and with (triangle). Closed symbols represent localities of checked rather blunt tips; 16 scale rows around mid-body; 14 specimens, open symbols represent unchecked literature records; sources: Blanc & Fretey, 2000; Broadley, 1998; scale rows in the neck region; 14 scale rows anterior to Lawson & Ustach, 2000. vent; 160 ventrals, slightly keeled; 58 subcaudals. Coloration. Dorsal coloration in life is bronze; sev­

Description. Slender tree viper, 480 mm total length, eral scale tips and keels being dark-brown, forming an tail 95 mm; very short and sturdy head, IODISED 2.3; irregular pattern of broken crossbars; ventrally cream­ rostral three times as wide as deep, surmounted by six white to yellowish; iris yellow (Fig. 3). In preservative suprarostrals; viewed from the front the three median (3 % formaldehyde transferred to 70 % ethanol after two suprarostrals in the lower row increase in size from left months), colour changed to reddish-brown dorsally; to right; the upper row consists of three scales, the lat­ ventral scales turneduniform clear reddish brown, being eral ones larger than the median (Fig. 2c); five more intense in posterior third of body and tail, fading internasalsstrongly carinated; nasals undivided; a single anteriorly. scale row separates nasals from circumorbital scales; Etymology. The name refers to the hirsute appearance head scales heavily carinated and lanceolate; nine of the snake (!at. hirsutus =hairy, hirsute).

TABLE 1. Head-measurements and indices of specimens examined. Data forAt heris squamigera are based on all material examined (n=23). Damaged specimens were excluded from the analysis. Measurements in mm. For abbreviations see the material and methods section. HL HW IOD SED HL/HW IODISED HWIIOD Atheris hirsuta sp. nov. 14.3 10.7 6.9 3.0 1.3 2.3 1.6

Atheris squamigera Mean 20.2 15.0 8.1 4.8 1.4 1.7 1.8 Mode 16.4 11.1 8.2 5.3 1.1 1.6 1.4 SD 4.1 3.3 1.4 1.0 0.1 0.1 0.2 Min 12.5 8.5 5.3 3.3 1.1 1.4 1.4 Max 28.9 22.1 11.8 7.2 1.7 1.9 2.2

TABLE 2. Head scalation of specimens examined. Data for Atheris squamigera are based on the entire material examined (n=44). Damaged specimens were excluded from the analysis. For abbreviations see the material and methods section.

SRO INS IOS MTHS cos COS IOL ION SL SL IL IL PSL left right left right left right Atheris hirsuta sp. nov. 6.0 5.0 9.0 14.0 15.0 14.0 0.0 1.0 10.0 9.0 9.0 8.0 3.0

Atheris squamigera Mean 4 4 8 14 14 14 0 10 10 11 11 5 Mode 3 4 8 15 14 14 0 10 10 11 11 5 SD 1.5 0.6 0.7 1.1 1.4 1.3 0.3 0.4 1.0 0.9 1.0 1.2 0.6 Min 3 3 7 12 11 11 0 8 8 8 8 4 Max 8 6 10 17 17 17 2 12 12 12 13 7 A NEW ATHERIS SPECIES 57

TABLE 3. Body scalation and SYL in mm of specimens temperature are up to I 0°C. Mean annual temperature is examined. Data fo r Atheris squamigera are based on all about 25°C. Humidity fluctuates from 85% during the material examined (n=44). Damaged specimens were day to 90-100% during the night. This area is situated excluded from the analysis. • = + anal. For abbreviations see the material and methods section. within the equatorial climate zone, which is influenced by the southernpassat (Riezebos, Vooren & Guillaumet, MSR Ventralsa Sub- SVL 1994). Floristically it belongs to the -Congo Re­ caudals gion (Guillaumet, 1967). The holotype was found after Atheris heavy rain, on a dirt road between the small town ofTa"i hirs uta sp. nov. 16.0 160 58.0 480 and the SRET. This area is characterized by secondary rain forest. Atheris Behavioural remark. Compared to the sympatric A. squamigera chlorechis, A. hirsuta seemed to be more aggressive. Mean 19 154 52 428 When disturbed, it immediately coiled the anterior third Mode 20 158 55 450.0 of the body in an S-shape, ready to strike. SD 1.6 7.7 5.7 93.3 Distribution. So far only known from the type local­ Min 16 140 31 220 ity (Fig. 1 ). Max 22 171 65 605 DIFFERENTIAL DIAGNOSIS AND DISCUSSION Habitat. The TNP (6°10'-5°10'N, 7°20'-6°50'W) is Atheris hirs uta sp. nov. resembles A. squamigera in the largest protected rain forest area in . appearance but can be distinguished from the latter by Annual precipitation reaches 2200 mm in the south-west several morphometric, and morphological characters. and 1700 in the north-east of the park. Precipitation is Its most intriguing diffe rence is the elongated dorsal highest fromApril-May to June-July and from Septem­ scalation, especially on the head and anterior body, ber to October-November. A firstdry period lasts from which could not be observed to such an extent in any December to February. A second dry period normally specimens of A. squamigera. Further distinguishing occurs in August. Daily mean temperature varies be­ characters were: the number and arrangement of tween 20°C and 30°C; diurnal fluctuations in suprarostrals (SRO) (six in A. hirsuta; three or up to eight in A. squamigera, but usually odd numbers occur); the number of pairs ofsublinguals (PSL) (three pairs in A. hirsuta; four to seven in A. squamigera); and the shape of the head and neck scales. Scales in A. squamigera are never as strongly bent as in A. hirsuta (Fig. 2 d, e), which in this respect appears similar to A. hispida and A. acuminata. Atheris hirsuta has a much shorter snout and larger eye than A. squamigera, giving the head a more blunt and stocky appearance. This is expres;ed by the IODISED index, which is 2.3 in A.

c

FIG. 2. Lateral (a) and ventral (b) aspect ofAt heris hirsuta sp. nov. (holotype, SMNS 1 1333) fromTa" i-National Park, Ivory Coast; scalation of the snout (c); dorsal scales from Atheris FIG. 3. Atheris hirsuta sp. nov. (holotype, SMNS.113 33) hirsuta (d) and A. squamigera (e) (ZSM 375 I 1909). from Ta"i National Park, Ivory Coast. 58 R. ERNST ET AL.

hirsuta, compared with a maximum of 1.9 (mean 1.7) in pattern, also used to distinguish it from the closely re­ A. squamigera. The number of mid-dorsal scale rows lated A. squamigera (Lawson, 1999). Two distinctive (MSR) in the holotype of A. hirsuta is comparatively East African species, A. hispida and A. acuminata, also low - only 16, which equals the lowest counts found in posses acuminate or lanceolate dorsal scales, but differ A. squamigera (SMNS 4213; ZMB 20481). fromA. hirsuta by having fewer transverse head scales Atheris hirsuta differed from the sympatric A. and a lower number of suprarostrals, as well as fewer chlorechis by the number of suprarostrals (7-8 in A. pairs of sub linguals. In addition, A. hisp ida has a higher chlorechis, arranged in a lower row of four and an upper number of interocunasals. Atheris acuminata has a row of three to four scales) and the maximum transverse lower number of circumorbital scales, as well as lower

head scale count (MTHS) between the posterior numbers of supralabials. For a summary of differences supralabials (SL) (25-27 in A. chlorechis; 14 in A. between A. hirsuta and other A theris species, see Ta­ hirsuta). has higher infralabial ble 4. counts (IL) (I0-1 1; 8-9 in A. hirsuta) and higher num­ Several authors pointed out that the high variability bers of dorsal scale rows at mid-body (25-37). Scales of within the genus Atheris, and especially A. squamigera, A. chlorechis are not as strongly keeled and the keel may at least partially be due to incorrectly combining only stretches across two thirds of the entire scale, leav­ distinct species within a single taxon (Lawson, 1999; ing a smooth, slightly depressed area uncovered. Our Hughes pers.comm.). For example, A. hisp ida had long SEM-photos of A. hirsuta revealed no serration of keels been taken as a of A. squamigera (Laurent, on lateral scales, known to occur in some A. chlorechis 1955). The same is true forA. broadleyi, which had been (Groombridge, 1980, cited by Broadley, 1996, 1998). referred to as a colour morph of A. squamigera (Perret Atheris hirs uta differs from the East Africanspecies & Mertens, 1957; Broadley, 1998), until its description A. nitschei, A. rungweensis, A. desaixi, A. katangensis by Lawson ( 1999). The resurrection of the species status and A. ceratophora by having fewer transverse head of A. subocularis has recently been published (Lawson scales, fewer interocunasals (ION) and higher numbers et al. 2001). We herein adopted the recognition of A. of dorsal scale rows at mid-body. Dorsal scalation in anisolepis as junior synonym of A. squamigera by these species is never as elongate as in A. hirsuta. The Lawson & Ustach (2000). However, these authors do latter lacks serrated keels on lateral scales, present in the not consider the difference in suprarostral scale counts species mentioned above. The lack of elongate (seven to eight in A. anisolepis, three to seven in A. supraoculars that formhorn-like projections above the squamigera), mentioned in Broadley ( 1998). We found eyes furthermore distinguishes A. hirsuta from A. this character to be consistent in four museum speci­ ceratophora. Keels in A. rungweensis and A. desaixi mens (SMNS 8361; ZMB 28987; ZSM 275/1996; ZSM end before the scale tip, whereas they reach the tip in the 375/1909;) and an additional specimen (ZMUC other members of this group, as well as in A. hirsuta. A. R68269) previously recognized as A. anisolepis rungweensis and A. desaixi have more interoculabials (Broadley, 1998). Referring to data presented by (IOL), as well as higher numbers of infralabials. The Broadley (1998), Hughes (pers. comm.) has pointed out number of pairs of sub linguals is higher in A. desaixi. the great range in ventral counts (males: 133-169, varia­ Atheris broadleyi from Cameroon is distinguished from tion: 36 scales; females: 141-175, variation: 34 scales) A. hirsuta by having more interoculabials, a higher in A. squamigera. In his opinion, a variation of more number of scale rows at mid-body and fewer than 30 in ventral scale counts within one sex is sugges­ interorbitals (IOS), and by its consistent dorsal colour tive of more than one species being involved.

TABLE 4. Comparison of characteristics among Atheris hirsuta sp. nov. and other Atheris species. Differences to Atheris hirsuta are marked bold. SLS = serrated keels on lateral scales; ESO = elongate supraoculars; * = no data; + = present; - = absent. For further abbreviations see the material and methods section. Data referring to Atheris squamigera represent mean values of specimens examined. Data referring to species other than Atheris hirsuta and Atheris squamigera adapted from: Broadley, 1998 and Lawson, 1999.

SRO INS IOS MTHS cos IOL ION SL IL MSR PSL SLS ESO hirsuta 6 5 9 14 14-15 0 9-10 8-9 16 3 squamigera 4 4 8 14 14 0 10 11 19 5 nitschei 3-7 4-5 6-12 18-20 10-17 0-2 2-5 8-13 9-15 23-34 3-6 + rungweensis 3-7 5-6 9-13 24-26 15-18 1-2 3-4 9-12 11-13 23-33 2-3 + desaixi 6-7 5 8-1 1 22 14-17 1-2 2-3 10-12 11-14 21-31 4-6 + ceratophora 5-9 4-5 7-11 19-20 13-19 0-1 2-4 7-1 1 8-12 19-27 1-3 + + katangensis 3-6 5-6 9-1 1 20-22 14-17 0-1 2-3 9-12 11 23-31 3 + chlorechis 7-8 5 8-14 25-27 15-20 0-2 3-4 9-12 10-1 1 25-37 1-2 + broadleyi 3-7 3-5 3-8 * 12-16 0 3 9-12 9-12 17-23 * hispida 3 4-6 6-10 12 9-15 0 2 9-10 8-10 15-19 1-2 acuminata 2 3 5 10 11-12 0 6 7-8 14 1 A NEW ATHERIS SPECIES 59

Characteristic threat displays involving stridulation, From their experience in other regions (Rode!, Grabow, that have been reported for A. desaixi (Ashe, 1968) and Bockheler & Mahsberg, 1995; Rode!, Kouadio & erroneously forA. nitschei (Goetz, 1975), were not ob­ Mahsberg, 1999) they suggest that approximately only served when handling A. hirsuta. The dorsal scales of A. two thirds of the snake fauna ofTNP has been found so hirsuta showed no micro-ornamentation, as has been il­ far. With this description the known species score of lustrated by Price ( 1982, Fig. 6a) and Groombridge TNP rises to 40. ( 1980, Figure 184, cited in: Broadley, 1996) for A. squamigera. In our opinion, it remains questionable ACKNOWLEDGEMENTS whether this is really a fe ature of diagnostic value. MOR was supported by a post-doctoral scholarship Among the material examined in SEM-analyses, we from theGerman Academic Exchange Service (DAAD). found A. squamigera showing such ornamentation as The following colleagues helped in sending specimens well as specimens showing no ornamentationat all. from their collections: A. Schltiter (SMNS), W. Bohme Atheris hirs uta clearly differs from thefrequently ob­ (ZFMK), J. Hallermann (ZMH), J. Rasmussen served colour patterns in A. squamigera. The latter are (ZMUC), F. Glaw (ZSM) and R. Gunther (ZMB). most often apple green to turquoise blue with yellow TROPENBOS-C6te d'Ivoire helped with transporta­ crossbands, occasionally uniform spectrum yellow tion and various administrative services. Lodging (Lawson, 1993) or violet (Lawson, 1999), or a mottled facilities in TNP were provided by the "Centre de Re­ combination of these colours, and sometimes light cherche en Ecologie" and the "Projet Autonome pour la brown (this paper). Conservation du Pare National de Tai." Working in Another strong argument forthe distinctiveness of A. TNP was kindly permitted by the Commandant K. hirs uta from A. squamigera is their allopatric distribu­ N'Dri. The "Projet Autonome pour la Conservation du tion. The reports of the latter species from Senegal Pare National de Ta'i'',and the "Ta·i Monkey Project" (Lawson, 1999) and the Ivory Coast (Spawls & Branch, provided logistic support. Research permission was 1995; McDiarmid, Campbell & Toure, 1999) are not given by the "Ministere de l'Enseignement Superieur et substantiated by voucher specimens and are most prob­ de la Recherche Scientifique," of the Republic oflvory ably repetitions of earlier misidentifications in the Coast. The access permit to TNP was issued by the literature. The westernmost verified A. squamigera "Ministere de la Construction et de !'Environnement." record is from "" (Werner, 1897; Sternfeld, 1910), SEM-analyses were done with the assistance of Dr. R. based on ZMB 13777. There were no precise locality Wolf. D. Broadley, B. Hughes and W. Wiister provided data for Buettner's specimens, but they probably came valuable comments on a previous draft of the manu­ fromthe forestedTogo hills, which now lie on the bor­ script. der between Togo and , since fo rmer western Togo now constitutes the Volta Region of Ghana, ac­ REFERENCES counting forthe inclusion of Ghana within the range of this species by Leeson ( 1950) and Hughes & Barry Ashe, J. ( 1968). A new bush viper. J. £. Afr . nat. hist. Soc. (1969). Werner (1897) provided data �o r four natn. Mus. 27, 53-59. uncatalogued specimens of A. squamigera with low Blanc, C. P. & Fretey, T. (2000). Les de la reserve counts for midbody scale rows (17-19), and a head de f;une de la Lope et de la f6ret des abeilles (). which he assigned to the East African species A. Bull. Soc. zoo!. Fr. 125, 28 1 -292. ceratophora. However, Sternfeld( 1910) only listed two Broadley, D. G. (1996). A review of the tribe Atherini A. squamigera from Togo, collected by Buettner, with (Serpentes: Yiperidae), with the description of two the Berlin Museum number 13777, but claimed that the new genera. Afr . J. Herpetol. 45, 40-48. species was new for Togo. This "Togo" population is Broadley, D. G. (1998). A review of the genus Atheris separated from populations in south eastern east Cope (Serpentes: Yiperidae), with the description of a to by the "Dahomey Gap", an area well known to new species from Uganda. Herpetol. J. 8, 1 17-135. form a zoogeographic barrier for forest species (e.g. Goetz, J. C. (1975). The vipers of Africa south of the Schi0tz, 1967). This area is occupied by "forest-savanna Sahara. Bull. New York herp etol. Soc. 12, 3-11. mosaic", with only small patches of forest remaining, Groombridge, B.C. ( 1980). A phyletic analysis of viperine . Unpub. PhD thesis, Council for National and has existed in its present form for some 10 OOO years. The present situation may be the cumulative ef­ Academic Awards, UK. fe ct of successive Pleistocene dry periods, due to Guillaumet, J. - L. ( 1967). Recherches sur la vegetation et la flore de la region du Bas-Cavally (Cote d'Ivoire). climatic oscillations (Moreau, 1963, 1969; Jahns et al., 1999). Memoires Offi ce de la Recherche Scientifi que et Te chnique Outre-Mer. 1-247 + 15 plates. Although our description of A. hirsuta is based on a 20, Hughes, B. & Barry, D. H. ( 1969). The snakes of Ghana: a single specimen, we believe the differences to be suffi­ cient to justify its description at the species level, checklist and key. Bull. Inst. fo nd. Afr . noire ser. A, 31, 1004-1041. thereby making the name accessible forfurther studies M. of the snake fauna of the Ivory Coast. Rode! & Jacobi, (2001). The African vipers of the Atherini Mahsberg (2000) recorded 39 snake species from TNP. complex. http://www.kingsnake.com/atheris/. 60 R. ERNST ET Al.

Jahns, S., Hiils, M. & Sarnthein, M. ( 1999). Vegetation Perret, J.-L., & Mertens, R. ( 1957). Etude d 'une collection and climate history of west equatorial Africa based on herpetologique faite au Cameroun de 1952 a 1955. a marine pollen record off (site GIK 16776) Bull. Inst. fo nd. Afr . noire ser. A 19, 548-60 I. covering the last 400,000 years. Rev. Palaeobot. Price, R. M. (1982). Dorsal snake scale Palynol. 102, 277-288. microdermatoglyphics: ecological indicator or Laurent, R. F. (1955). Diagnoses preliminaries de taxonomic tool? J. Herpetol. 16, 294-306. quelques Serpents venimeux. Rev. Zoo!. Bot. Afr . 51, Riezebos, E. P., Vooren, A. P. & Guillaume!, J. L. (1994). 127- 139. Le Pare National de Tai', Cote d 'lvoire. Tropenbos Lawson, D. P. ( 1993 ). The reptiles and of Series 8, Wageningen. Korup National Park Proj ect, Cameroon. Rode!, M.-0., Grabow, K., Bockheler, C. & Mahsberg, D. Herpetological Natural History. 1, 27-90. ( 1995). Die Schlangen des Comoe-Nationalparks, Lawson, D. P. ( 1999). A new species of arboreal viper Elfenbeinkiiste (Reptilia: : Serpentes). (Serpentes: Viperidae: Atheris) from Cameroon, Stuttgarter Beitr. Naturk., Ser. A. 528, 1-18. Africa. Proc. Biol. Soc. Washington. 112, 793-803. Rode!, M.-0., Kouadio, K. & Mahsberg, D. ( 1999). Die Lawson, D. P. & Ustach, P. C. (2000). A redescription of Schlangenfauna des Comoe-National parks, Atheris squamigera (Serpentes: Viperidae) with Elfenbeinkiiste: Erganzungen und Ausblicke. comments on the validity of A. anisolepis. J. Herpetol. Salamandra. 35, 165-180. 34, 386-389. Rode!, M.-0. & Mahsberg, D. (2000). Vorliiufige Liste Lawson, D. P., Noonan, B. P. & Ustach, P. C. (200 1). der Schlangen des Tai-Nationalparks/Elfenbeinkiiste Atheris subocularis (Serpentes: Viperidae) Revisited: und angrenzender Gebiete. Salamandra. 36, 25-38. Molecular and Morphological Evidence for the Schi0tz, A. ( 1967). The Treefrogs (Rhacophoridae) of Resurrection of an Enigmatic Taxon. Copeia 2001, West Africa. Sp olia zoo!. Mus. haun. . 25, 1-346. 737-744. Spawls, S. & Branch, W. R. (1995). Th e dangerous snakes Leeson, F. ( 1950). Identification of snakes of the Gold of Africa. Halfway House: Southern Book Publishers. Coast. The Crown Agents for the Colonies, London. Sternfeld, R. ( 1910). Die Schlangenfauna . Mitt/. McDiarmid, R. W., Campbell, J. A. & Toure, T. A. ( 1999). Zool.Mus. Berlin 4, 207-236. Snake Sp ecies of the world. A taxonomic and Werner, F. (1897). Ueber Reptilien und Batrachier aus geographic reference. 1, Herpetologists' League, Togoland, Kamerun und Tunis aus dem Kgl. Museum Washington, D.C.. fiir Naturkunde in Berlin. Verh. zool.-bot. Ges. Moreau, R. E. ( 1963 ). Vicissitudes of the African biomes Wien.47, 395-408. in the late Pleistocene. Proc. zoo/. Soc. London 141, 395-421. Moreau, R. E. ( 1969). Climatic change and the distribution of forest vertebrates in West Africa. J. Zoo/., lond. Accepted: 14. 1. 02 158, 39-61.

APPENDIX Neu-Kamerun; ZMB 30690, Aj osh6he am Nyong; ZMB COMPARATIVE MATERIAL EXAMINED: 30719, Aj oshohe am Nyong; SMNS 4213, Acra, Kamerun; SMNS 4264, Malimba, W.Afrika; Atheris squamigera ZFMK15619, Kamerun & FernandoP6, 1951154 1963, BURUNDI:ZSM 275/1996, E:VIII.93-VI.96, Burundi, Kumba; 1951/54 1963, Moka /Bioko; ZFMK5452-56, Musigati; Kamerun+Fernando P6, 1951/54 1963, Moka /Bioko; CAMEROON: ZMH R063 l 6 old no. 1760, Cameroon; ZFMK 15618, Kamerun &FernandoPo, 1951/54 1963, ZMH R0632 l old no. 1861, Cameroon; ZMH R063 l l Nyasoso/Kupe; old no. 4973, Jaundebezirk (Yaounde) an der CONGO: ZMUC R68269 (800), Menengue, 4°16'S- Kribistrasse, S Yaounde, Cameroon; ZMH R06323 old 1l047'E, Congo; ZMUC R68270 no. 4493, Esosung, Bakossi Mountains, Bezirk Johann­ (RJD 12),Tchissanga, 4°32' S- 1l046'E, Congo; ZSM Albrechts-Hohe (=Kumba) 1060 m a.s.l., Cameroon, 130-137/1999 V.1995, Brazzaville umgeb.; Africa; ZMH R063 13 old no. 4949, Kuti, BarnumCoun­ DEMOCRATIC REPUBLIC OF CONGO (ZAIRE): try, 1040 masl, Cameroon; ZMH R06328 old no. 5131, ZMH R063 l 7 old 124, Belgisch Kongo, Democratic Central AfricaMolundu; ZSM 375/1909a, Dibongo bei Republic of Congo; SMNS 8361, Zaire, Afrika, Abdea Kamerun; ZSM 375/1909b, Dibongo bei Abdea Prov.Kivu Station Irangi; ZMB 37709, Leopoldville; Kamerun; ZSM 9611978, Esosung, Bakossi Mountains, ZMB 37807, Leopoldville; ZMH R06326 old 184, Kamerun; ZMB 20342, Douala, Yossplatte; ZMB Leopoldville: Kinshasa; 2048 1, Douala, Yossplatte; ZMB 15887-88, GABON: ZMH R06320 old 1098,Gabon; Joh.Albrechtshohe; ZMB 21727, Longji, Kamerun; TOGO: ZMB 13777,Togo. ZMB 28987, Longji, Kamerun; ZMB 24302, Nj ong, A NEW ATHER!S SPECIES 61

Gazetteer (only localities of West and Central African CONGO: Leketi, 2°34'S 14°17'E; Menengue, 4°16'S distributional range of A. squamigera included): 11°47'E; Tchissanga, 4°32,S 1 1°46'E; Lukolela, 5°23'S CAMEROON: 34 km N of Lolodorf, 3°14'N 10°44'E; 24°32'E; Prov.Kivu Station Irangi, 2°30'N 28°0'E; Nj ong, 3°17'N 9°54'E; Bitye, 3°l'N 12°22'E; Metet, Brazzaville umgeb., 4°16'S 15°17'E; 3°26'N 11°45E; Dibongo bei Abdea 3°47'N 10°6'E; DEMOCRATIC REPUBLIC OF CONGO (ZAIRE): Jaundebezirk (Yaounde) an der Kribistrasse, S Banana, 6°l'S 12°24'E; Leopoldville: Kinshasa, 4° 1 8'S Yaounde, 3°52'N 11°31 'E; Longji, 3°5'N 9°58'E; 15°18'E; Leopoldville,4°19'S 15°13'E; Avakubi, Boumir Camp, Dja Forest Reserve, 3°9'N 13°0'E; Kinshasa, 1°20'N 27°34'E; Niapu, Kinshasa, 2°25'N Aj oshohe am Nyong, 4°0'N 13°34'E; Fernando P6, 26°28'E; Medje, Kinshasa, 2°25'N 27°18'E; Akenge, Kumba, 4°43'N 9°1 l'E; Southwest Province, 5°25'N Kinshasa, 2°56'N 26°50'E; Rungu, Kinshasa, 3°1 l'N 9°20'E; Esosung, Bakossi Mountains, Bezirk Johann­ 27°52'E; Mbanza-Ngungu, Kinshasa, 5°15'S 14°52'E; Albrechts-Hohe (=Kumba) 1060 m a.s.l. Africa, 5°59'N Kinsuka, Kinshasa, 5°15'S 15°13'E; 14°26'E; Molundu, 2°2'N 15°13'E; Douala, Yossplatte, GABON: Makande, 0°47'S 11°58'E; 4°3'N 9°42'E; IVORY COAST: Tai NP, 5°50'N 7°19'W; TOGO: Togo, no precise locality available.